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Bruce K. Holst1, David Amaya2, Ella Baron2, Marvin Paredes2, Elma Kay3 1Marie Selby Botanical Gardens, 2 Ian Anderson’s Caves Branch Botanical Garden, 3University of Belize

© Marie Selby Botanical Gardens ([email protected]), Ian Anderson’s Caves Branch Botanical Garden ([email protected]). Photos by D. Amaya (DA), E. Baron (EB), W. Collier (WC), B. Holst (BH); J. Meerman (JM), L. Munsey (LM), M. Paredes (MP), P. Nelson (PN), D. Troxell (DT) Support from the Marie Selby Botanical Gardens, Ian Anderson’s Caves Branch Jungle Lodge, Environmental Resource Institute - University of Belize [fieldguides.fieldmuseum.org] [964] version 1 11/2017

The Tillandsia in Belize includes approximately 30 , which can be found growing singly, or in large colonies, and can usually be identified by their non-spiny leaves, often flattened branches, symmetrical , free petals, and often colorful flower clusters which fade quickly after flowering. They are most always epiphytic growing on and shrubs to gain better access to sunlight; an occasional species is found on rocks or on the ground (e.g., T. dasyliriifolia). Many have gray/silvery, scaly leaves (e.g., T. pruinosa, T. streptophylla). The scales (or “”) help capture water and nutrients from the environment. Some spe- cies form water-holding tanks by means of their overlapping leaves. These tanks are rich with nutrients from the environment, provide sustenance for the , and create important habitat for animals in the forest canopy. The genus is found throughout Belize, but reaches its peak of species diversity on the high summits of the Maya Mountains. Many species are adapted for pollination, with brightly-colored bracts and flowers. The seeds of all species in the genus develop inside leathery capsules, which split open into three parts when mature; they are topped by a fluffy coma (resembling a tuft of “hairs”) and then may be dispersed by the wind. Important distinguishing characteristics of the species are the shape of the rosette, the shape and degree of branching of the inflorescence, and the color and length of the various bracts and flowers. All species in Belize, except T. utriculata, produce one or more “pups” or “offsets” from the mother plant and continue to grow after the mother plant flowers and dies. This growth habit can eventually form extensive clonal colonies. District Abbreviations: Belize (B), Cayo (Ca), Corozal (Co), Orange Walk (OW), Stann Creek (SC), Toledo (T). Elevations are for Belize only. Photos included not taken of Belizean are identified in the text. Identification Guide to Tillandsia species of Belize Group 1. Stems elongate, leaves separated from one another along the stem; petals emerald green or yellow (2 species) Groups 2–7. Stems short, leaves tightly arranged in a rosette; petals purple, pink, greenish, or white (page 2) Tillandsia usneoides  Known from Ca, OW, T; 100–580 m elev.  The most distinctive til- landsia, but often over- looked; the pendent, gray foliage that doesn’t even look like a bromeliad is unique, and the emerald- green, fragrant flowers are rarely observed.  Known as . Stems can reach several meters The growth form of T. usneoides Capsules thin, slender, splitting Flowers small, green, often over- long and are soft (BH) (left) is similar to that of some into three parts as in all looked, but pleasantly fragrant (pho- lichens (pictured on right; EB) (photo of plant; BH) to of Florida plant; WC)  Common and widespread in Belize in B, Ca, Co, OW, T; 40–610 m elev.  Forms large colonies; distinct with tight “ball” shape, as well as the elongated stems, dense gray scales, reddish floral bracts, and yellow petals and anthers.

Plants exhibit clumping, branching Leaves spaced out along the slender, un- Flowers all yellow and protrude growth (EB) stout stem with heavy covering branched, with reddish/orangish above the floral bracts (EB) of grayish trichomes (EB) bracts and yellow flowers (PN)

Tillandsia (Bromeliaceae) of BELIZE 2

Group 2. Plants with bulbous bases, often clumping, often growing in multiple orientations in relation to their attachment (see also T. dasyliriifolia in group 6, with bulbous base, but with a large inflorescnece and long-spreading branches) (5 species) Groups 3–7. Plants not bulbous-based, the rosette form a narrow to broad vase (page 3)  Found in seasonally dry forests in all Districts; 5– 550 m elev.  Distinguished by the bulb- ous base, narrow, leathery, silvery green, spreading, re- curved, or twisting leaves; red to orange floral bracts, and purple corollas; plants often colonized by ants.

Distinct bulbous base, leaves Inflorescences narrow, elongate, Individual inflorescence Floral bracts sometimes infused spreading, often twisted (EB) simple, or more often compound; branches are few and held erect, with green and red, turning green scape bracts leaf-like (EB) and are flattened (EB) after flowering (PN)  Found in all Districts, and in many habitats except the wettest ones in the south; 0– 760 m elev.  Distinguished by the gray, bulbous base, few cylindric, narrow leaves, bright red bracts, lavender corollas.  Often found hanging upside down individually or in col- onies, and inhabited by ants. Medusa-like growth with bright Plants are found growing at Large, bulbous base, and con- Floral bracts red; corollas purple, bracts and flowers (WC) every angle, and often inhabited trasting leaf and sheath coloration; tubular, with protruding anthers by ants (BH) leaves are deeply channeled (BH) (MP)  Rare in Belize, known from a single collection from Lit- tle Quartz Ridge, T; 700– 750 m elev.  Similar to T. bulbosa and T. pseudobaileyi, but distin- guished by the prominent bands of spots on the leaf sheaths.  Images here taken of a spec- imen from . A poorly known species in terms As with other species in the pseudobulbous group, plants are found grow- Leaves distinct with bands of ma- of the herbarium and photographic ing at multiple angles, often downwards (EB) roon spots (EB) record (EB)  Discovered in Belize in 2017 by Caves Branch Bot. Gard. staff, from OW; 10– 20 m elev.  Rare in Belize; growth simi- lar to T. bulbosa, but leaves distinctly “lined” toward base and with dull gray col- oration, compared with not lined and with glossy, dark green leaves. Habit pictured here is of a plant in Leaves and bulbous bases T. pseudobaileyi on right, compared Flowers are similar to those of T. cultivation at Selby Gardens (PN) have fine vertical lines (DA) with T. bulbosa (left; DA) bulbosa (PN)

Tillandsia (Bromeliaceae) of BELIZE 3 Tillandsia ×jaguactalensis  Rare in Belize, only known from citrus fields in Ca; 80 m elev.  Similar to T. brachycaulos, but with elongated flower cluster, and to T. strepto- phylla, but flower cluster compact, versus broadly branching. This is a natural hybrid between these two species. To date only known from citrus The leaves are widely spreading Inflorescence bracts are pale pink Flowers are tubular, purple, and orchards in Belize (EB) and densely covered with silvery and elongated (EB) with protruding flower parts (EB) scales (WC)  Known from southern Be- lize in Ca, OW, SC, T; 130–750 m elev.  Distinctive compact plant with bulbous base, heavy covering of scales, few curved or twisted and deep- ly channeled leaves, pink floral bracts, and purple corollas.

Twisted leaves and compact form Even in vegetative state, distin- Inflorescences paddle-like, strongly Flowers with purple, tubular, purple with bulbous base; plants grow at guished by the heavy, scaly flattened, mostly shorter than the corollas, with protruding, yellow various angles from host (EB) nature, and few leaves (EB) leaves (PN) anthers (BH)  Most common in lowland areas, in B, Ca, Co, OW, SC, T; 0–600 m elev.  The most photographed tillandsia in Belize! A ro- bust species, often teaming with ants; distinguished by the large bulbous base, heavily scaly nature, twisted leaves, and multi-colored flower cluster. Attractive , seen here in Leaves are usually twisted and Plants can grow in almost any As with all species in this group, forests of Sarstoon-Temash warped, channeled and heavilly orientation; scape and primary corollas are purple, tubular, and National Park (BH) covered with trichomes (BH) bracts are pink to coral. (EB) with protruding flower parts (EB) Group 3: Leaves narrow at the base and along entire length, ≤ 3 mm wide (3 species) Groups 4–7: Leaves broad at the base, ≥ 5 mm wide, tapering evenly to a point, or broad and with parallel margins (page 4)  A rare species from Ca, SC, T; 160–750 m elev.  Distinguished by the narrow leaves and basal sheaths that are blackish and to- gether form a fluffy ball, and the delicate, greenish flower cluster and bracts, and lavender petals.

The smallest tillandsia in Belize; Leaves very fine and thin, and Inflorescence much longer than Petals and stamen filaments pale produces compact offsets and can pointing in every direction (EB) leaves, with greenish bracts (EB) lavender, the petals spreading occasionally branch (MP) widely and anthers protruding (EB)

Tillandsia (Bromeliaceae) of BELIZE 4  Where found, B, Ca, OW, SC, T, this species often occurs in dense colonies; 80–750 m elev.  One of the narrowest-leaved tillandsias, distinguished by the reddish bracts, laven- der/purple corollas, finely spotted leaves and floral bracts, and non- stoloniferous growth. Leaves narrow, much shorter Many plant parts covered with Lateral inflorescence branches slen- Corolla tubular, purple with than inflorescences; often grows fine reddish spots (EB) der, spreading from main axis (EB) protruding flower parts (EB) in large colonies (BH)  Found in B, Ca, OW, SC, T; 30–900 m elev.  Identified by the narrow leaves, compact flower clusters, and stoloniferous growth; foliage is often reddish in the sun; confused with T. festucoides, but has elongated stolons and short- er inflorescence branches (see above). Grows in large colonies; often Elongated stolons are often con- Inflorescences are approximately the Compact flower cluster has with reddish tinge to the narrow spicuous and help distinguish it same length as the leaves (EB) short lateral branches and leaves in the Pine Ridge (EB) from T. festucoides (EB) greenish to dull red bracts (PN) Group 4: Leaves strap-shaped, with mostly the same width along their length (3 species) Groups 5–7: Leaves narrowly triangular, narrowing gradually from the base to the tip (page 5) Tillandsia excelsa  Restricted to high eleva- tions in Belize in Ca, T; 700–1100 m elev.  Distinguished by the large, branched inflorescence with bright colors of orange, red, and yellow. The stiff, lateral inflorescence branches have long primary bracts at their bases and the leaves are spotted below. Sharon Matola of the Belize Young leaves are usually spot- Inflorescence erect, stiff and Petals are purple, and contrast with Zoo exhibits a “medium-sized” ted on the lower leaf surface, “spiky” with horizontal branches the red and green bracts (MP) bromeliad specimen (BH) and soft (JM) and red to orange-red bracts (BH)  One of the rarest species in Belize, found on the highest peaks of Ca, SC, T; 700– 1000 m elev.  A colorful, relatively small species, with reddish leaves (especially on the lower surface); inflorescences shortly branched, usually pendent, with long primary bracts; the petals purple. The plant is often with pen- The difference between upper Primary bracts red, recurved, as Corolla tubular and flares at the tip; dent inflorescences (PN) and lower leaf surface color is long as the inflorescence branches flowers parts are included within notable (DA) (PN) (MP)

Tillandsia (Bromeliaceae) of BELIZE 5  A rare species in Belize, restricted to high mountains of Ca, T; 800–1100 m elev.  Similar to T. anceps because of the paddle-shaped flower cluster, but with broader leaves, inflorescenses often more than 1, found in axils of leaves (versus terminal), and with bright orange-red bracts. Plants usually with 2 or more Inflorescence paddle-shaped, with Each flower is subtended, and Petals are purple and the protruding inflorescences at a time (BH) brightly colored scape and floral mostly covered by the orange-red anthers bright yellow (PN) bracts (LM) floral bracts (PN) Group 5: Leaves with colored stripes, bands (see also vittata in separate guide), or with purple-black leaf bases; pet- als purple, greenish, or white (4 species) Groups 6, 7: Leaves mostly of uniform color (unless in bloom); petals purple (page 6)  Found in humid forests of Ca, SC, T; 10–1000 m elev.  Distinguished by long, nar- row leaves with maroon stripes at the base; flat, pad- dle-like flower clusters with pink, green, or whitish floral bracts, and lavender flowers.

Leaves long, soft, narrow; flower Reddish vertical stripes usually Flower clusters broad, paddle- Petals spreading, lavender; anthers cluster pink, green or whitish (BH) present at the leaf bases (EB) like (PN) and stigma do not protrude beyond the petals (PN)  Found in lowland semide- ciduous forests in OW; 10 m elev.  One of the rarest, most dis- tinctive bromeliads in Be- lize, this was discovered for the first time in mid-2017 by Caves Branch Bot. Gard; it is distinguished by its twisted, gray-banded leaves, and pink inflorescences. Inflorescence greatly exceeds the Leaves are gray, twisted, and Inflorescence has long branches Flower bracts and corolla are pink, leaves and is widely branched with strong horizontal banding and widely spaced flowers (plant and anthers and stamens protruding (DA) (DA) from ; PN) (plant from Panama; PN)  A rare species in Belize, restricted to high mountains of Ca, T; 800–950 m elev.  Distinguished by the narrow leaves that are puple-black at the base, an inflorescence as long as the leaf blades, red to green bracts, and pur- ple corollas.

Tightly clumping form, narrow Leaf bases are distinctly dark Inflorescences elongate, arching Petals purple with white tips, an- leaves and dense colony (BH) purple-black (EB) or erect, with a combination of thers protruding; scape bracts reddish to greenish bracts (BH) reddish, floral bracts green (EB)

Tillandsia (Bromeliaceae) of BELIZE 6 Tillandsia orogenes  Another rare species in Be- lize, found only on the high- est peaks of the Maya Mountains, often growing on Colpothrinax cookii, in Ca, T; 980–1140 m elev.  Identified by narrow stiff leaves and dark bases, spreading bright red inflo- rescences that lave long primary bracts. Dense, clumping growth and elon- Leaves narrow, strict, with The inflorescences are much Primary bracts are longer than the gated inflorescences; shown here in blackish bases (BH) longer than the leaves, and the short lateral branches; petals hidden the Maya Mountains (BH) primary bracts are reflexed (LM) purple, anthers yellow (LM)

Group 6: Inflorescence > 2x beyond the leaves, branches elongate, slender and flowers widely spaced along them (3 species) Group 7: Inflorescence less than 2x beyond the leaves, if branched then with densely arranged flowers (page 7, below) Tillandsia dasyliriifolia  A species known from coastal and southern Belize and found in all Districts; 0–680 m elev.  Inflorescence similar to T. izabalensis, but plants dis- tinguished by the bulbous base, the flower cluster which produces “pups” after flowering, and floral bract length (<23.2 vs >23.4 mm). Base bulbous with large basal leaf In nature, the young leaves are Bromeliad “pups” develop along Floral bracts usually red and flowers sheaths; leaves strongly covered often pink to bronze (EB) the inflorescence branches (EB) greenish; inflorescence branches by silvery scales (PN) strongly “zig zag” (PN) Tillandsia izabalensis  A coastal species found in Ca, Co, SC, and T; at low elevations, 5–20 m.  The tallest tillandsia in the country, and belonging to a complex of several species  Often confused with T. dasyliriifolia. See under that species, as well as the close- ly related T. utriculata.

Exhibiting the large size of a Inflorescence few-branched Rosettes similar to T. utricula- Inflorescence similar to T. dasylirii- flowering specimen (EB) (compare with below; BH) ta (see below; DA) folia, but floral bracts longer (EB)  Moist forests, especially common in citrus groves of Ca, SC; 5–130 m elev.  A large plant distinguished by a spreading rosette, slen- der, straight inflorescence branches, and white petals. See similar species T. dasyliriifolia and T. izaba- lensis.

Rosettes are similar to T. izaba- Slender, straight inflorescence Leaves are stiff, light to dark Corolla tubular, twisted to one lensis, but inflorescences with branches can be green or burgun- green above, paler below; com- side; petals white; anthers and numerous, slender branches (EB) dy in color (EB) mon in citrus orchards (EB) stigma protruding (EB)

Tillandsia (Bromeliaceae) of BELIZE 7

Group 7: Inflorescence less than 2x beyond the leaves, if branched then with densely arranged flowers Group 8: Includes additional rare species and a note on hybrids (page 8, below)  This compact species grows in large colonies in the cit- rus orchards of Ca, and is also known from B, OW, T; 0–550 m elev.  Distinguished by flower cluster borne among the central leaves, and vivid red to coral color bracts; com- pare with T. variabilis, which has fewer leaves. Easily seen when in bloom, of- Often all parts of the plant are After flowering, the plants turn Purple flowers borne among the ten in citrus groves (EB) colored, and fade to green after green, even if in full sun; brown colorful bracts; anthers and flowering (EB) structures are fruits (EB) stigma protruded (EB)  Known from central to northern Belize in B, Ca, Co, OW; 10–400 m elev.  Recognized by the stiff, narrow, ascending leaves, stout flower clusters with few branches, reddish to green bracts, and laven- der/purple corollas. There are several color forms in Belize. Leaves silvery-gray, light to Colonial growth, with red and Multi-colored flower cluster, lat- An unusual form with greenish dark brown at base (BH) green colored inflorescences and eral branches short, stout. Com- floral bracts and white corollas (EB) numerous stiff leaves (BH) pare with “sp. 2” in Group 8 (EB) Tillandsia chlorophylla  A rare bromeliad in Belize, only known from “near the border of ,” T; elev. unknown.  Distinguished by the color- ful, compound inflores- cences with short branches and long primary bracts that envelop the short branches; leaves are slender and long.

Collected only once in Belize along the Guatemala border by William Primary bracts that subtend the short Purple petals protrude well beyond A. Schipp in the 1934 during the British -Guatemala Bounda- branches are as long as, or longer the bracts, and anthers and stigma ry Survey. The plant pictured here is from Guatemala and cultivated at than the branches (PN) beyond them (PN) Selby Gardens (PN) Tillandsia maya  Uncommon in Belize, and found only in the citrus or- chards of Ca; 80 m elev.  Intermediate between T. balbisiana, T. polystachia, and T. brachycaulos. The plant is less bulbous at the base than T. balbisiana, and the inflorescence is more elongated than T. brachy- caulos. It may be a hybrid. Discovered in Belize in 2014 Foliage similar to T. polystachia, Flower cluster bracts are red Flowers are erect, with purple by staff of Caves Branch Bot. see below (EB) and spread widely as in T. bal- petals, and with protruding flower Gard. (EB) bisiana (EB) parts (EB)

Tillandsia (Bromeliaceae) of BELIZE 8  Uncommon in Belize, though can be abundant in citrus orchards, Ca; 80–550 m elev.  Small plant with narrow, stiff, glossy green leaves, green to reddish inflo- rescence, and purple flow- ers; leaves can turn bronzy in the sun. Possibly hybridiz- ing, making ID difficult.

Relatively few leaves arching Inflorescence erect, elongated and Leaves are stiff, smooth, recurved Floral bracts often green with some to recurved; often found in slender, with leaf-like scape bracts or reflexed, and matte green (EB) red tinges; purple corollas tubular, Cayo citrus fields (MP) and short lateral branches (BH) and with protruding anthers (EB)  Found in broadleaf forests and citrus orchards of B, Ca, OW, SC, T; 25–610 m elev.  The relatively small size of the rosette, narrow leaves, simple or few-branched pink to red inflorescences, strong- ly flattened inflorescence branches, help distinguish it from other species.

Leaves are usually green in the Comparing inflorescences of T. When inflorescence is branched, Inflorescence branches flattened shade, reddish in the sun; the polystachia (left) and T. varia- the branches have long axes laterally; corollas tubular, purple, inflorescences are red during bilis; note lack of lengthy scape (compare with short branch axes with protruding stamens and stigma flowering (PN) bracts in T. variabilis (EB) in T. polystachia; EB) (EB)

Group 8: Additional rare or unknown species  There are three additional species documented in Be- lize that are poorly known from the herbarium and photographic record.  One is identified to species level, two are not.  There are likely more bro- meliad species to be discov- ered in remote areas of the country.

 Tillandsia sp. 1  Tillandsia sp. 2  Epiphyte on Maya Moun-  Lithophyte on Pine Ridge,  Epiphyte from CA, OW; 400–500 m tains, T; 750 m elev. Ca; 620 m elev.  This species belongs to the Tillandsia fasciculata complex,  Only known photo of a Be-  Leaves with coppery tinge but more study is needed to determine the correct name. lizean plant is the one shown above, grayish below.  Differs from typical T. fasciculata by having a broad, paddle- above of a plant in fruit.  Possibly an undescribed shaped inflorescence 4–5 cm wide (vs. 1–3 cm); both species  Bears a stout, simple inflo- species, it has the inflo- can have simple or compound inflorescences and the vegeta- rescence and according to the rescence simple, arching; tive portion of the plants is identical (EB) liteature, orange and green corollas purple with protrud- bracts when in flower (WC) ing stamens and stigma (EB)

Tillandsia (Bromeliaceae) of BELIZE 9

Illustrated Glossary

Letters in parentheses below refer to those in the illustrations.

 Floral bract (F): the modified leaf subtending a flower, which can be longer than, and obscure the calyx from view.  Flower (J): consisting of the sepals (together called calyx), the petals together called corolla), the androecium (the male part of the flower, or stamen, consisting of the filament and the anther), and the gynoecium (the female part of the flower, or pistil, consisting of the ovary, style, and stigma).  Flower cluster (C): the portion of the inflorescence consisting of the flowering region of the inflorescence, including the associated pri- mary and floral bracts, the axes bearing flowers, and the flowers themselves.  Habit (A). General shape and growth form of a plant.  Inflorescence (B): the flowering portion of the plant, which consists of the scape and flower cluster.  Inflorescence type: bromeliad inflores- cences can be simple (unbranched; see left-side of adjacent illustration, or T. an- ceps) or compound (branched; see right- side of adjacent llustration, or T. chloro- phylla, below). When compound, the branches can be short (e.g., T. polysta- chia), or elongate and spreading (e.g., T. utriculata).  Leaf (D): The vegetative portion of the plant, including the broad basal leaf sheath (Da) and the usually narrower blade, or lamina (Db).  Primary bract (G): the modified leaf at the base of an inflorescence branch; it can be colorful (e.g., T. leiboldiana), or small and green, and inconspicuous (e.g., T. filifolia)  Scale (see “” below).  Scape (I): the stalk that connects the vegetative portion of the plant to the flower cluster; the scape may be short and hidden among the leaves and bracts (e.g., T. brachycaulos), or elongate and evident (e.g., T. izabalensis). Note, the term “peduncle” is used for the same structure in many other plant families.  Scape bract (H): the modified leaf borne along the nodes of the scape that can be from scale-like to leaf-like in ap- pearance.  Trichome (E): minute structures analo- gous to plant “hairs” and often called scales, that cover the leaves of many bromeliads (seen as a gray cover on many tillandsias). Trichomes in bromeli- ads are often scale-like and have an ele- gant “mosaic-window” appearance. They help to facilitate the movement of water Tillandsia butzii. A: Habit, showing “bulbous” base and ramet (offset, or “pup”) devel- and nutrients into the plant, as well as to opment. B: Inflorescence. C: Flower cluster. D: Leaf; Da: leaf sheath; Db: leaf blade. E: help regulate water loss. Trichomes; Ea: Trichome from abaxial edge of leaf-sheath; Eb: Trichome from abaxial surface of leaf blade. F: Floral bract. G: Primary bract. H: Scape bract. I: Scape. J: Flower with floral bract. K: Anterior . L: Posterior sepals. M: Petal. N: Stamens. O: Ovary, style, and stigma. P: Stigma (enlarged).

Tillandsia (Bromeliaceae) of BELIZE 10

Tillandsia chlorophylla. A: Habit. B: Inflorescence (compound, or “branched”). C: Flower cluster. D: Leaf; Da: leaf sheath; Db: leaf blade. E: Trichome. F: Floral bract. G: Primary bract. H: Scape bract. I: Scape. J: Flower with floral bract. K: Anterior sepal. L: Posterior sepals. M: Petal. N: Ovary, style, and stigma. O: Stamens. P: Stigma (enlarged).

Note on Natural Hybrids Possible hybrids can be observed in several areas of Belize, particularly in the citrus orchards where many species thrive closely togeth- er. A more detailed study is needed to determine the possible parents of some hybrids, but we believe the following species may be in- volved in natural hybridization: T. balbisiana, T. brachycaulos, T. fasciculata, T. polystachia, T. streptophylla, and T. variabilis.

We thank Germán Carnevali, Ivón Ramírez, Juan Pablo Pinzón, and Brian Sidoti for their helpful comments, and Sue Sill for providing the glossary illustrations.